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Physicochemical and chemical properties of soils and BioC
The physicochemical and chemical characteristics of the soils and BioC, as well as selected chemical properties of the HAs isolated from the soil and BioC are shown in Table 1.
Table 1 Physicochemical and chemical characteristics of soils, BioC and isolated HAs.
Full size table

The properties of soils and BioC, such as the d, Corg, A, pH, and Q, were presented in detail previously4. Briefly, soils were characterised by a typical d value for mineral soils ≈ 2.60 g cm−3, and by a relatively low content of Corg and a high content of A. The pH of the soils was weakly acidic. The examined soils were characterised by low Q values, indicating a low content of organic structures dissociating to the negative surface charge (mainly carboxylic and phenolic groups). The HAs obtained from fallow and grassland were characterised by high QHA values (about 50 times higher in comparison with the Q values of fallow and grassland). The d value of BioC was typical for organic materials (1.46 g cm−3), moreover, the BioC contained a high content of OM, which was expressed as Corg. The pH of BioC was alkaline. This material was also characterised by a high Q value, which indicated its favourable sorption properties.
The results of our studies showed that the E2/6 values were similar for the HAs originated from the two studied soils, suggesting a similar ratio of lignin-type compounds resistant to humification to the structures with a high humification degree. The ΔlgK reached values of 0.83 and 0.86 for HAs isolated from grassland and fallow, respectively, indicating a low degree of HA humification (Kumada’s classification for low humification degree of HAs: ΔlgK = 0.8–1.1)33. Slightly higher ΔlgK values obtained for the grassland HAs compared with the fallow suggested a higher content of less humified compounds, such as cellulose, hemicellulose, and lignin34.
The ΔlgK of HAs isolated from BioC reached a value of 0.54, suggesting the presence of highly humified compounds, in comparison with soil HAs (Kumada’s classification for high humification degree of HAs: ΔlgK  8.0, above which the OH groups are deprotonated26, therefore we only report results in this pH range. Changes in the QHA values as a function of pH (Fig. 4A–D) were monotonic; these values increased towards an alkaline pH, which resulted from the fact that other fractions of functional groups dissociated successively at increasing pH values. Generally, in the first month of the experiment, the highest QHA values were observed for HAs obtained from fallow and grassland with the lowest BioC dose (Fig. 4A,C). This fact indicated that these HAs had the best sorption properties. In the last month of the experiment, the QHA values changed in an ambiguous way. The QHA at pH 9.0 values of HAs isolated from pure BioC were lower than those obtained from the soil, and moreover, BioC did not have an obvious effect on the QHA values of the soil HAs. Previous studies4 on impact of BioC on the physicochemical properties of Haplic Luvisol under different land uses, showed that BioC added to soil caused a significant increase in Q values in the last year of the experiment. Thus, we can conclude that BioC introduced OM with a variable surface charge but did not affect the soil’s QHA. It is possible that the BioC doses used in our experiment were insufficient to raise the QHA values.
Figure 4

Dependence of surface negative charge (QHA) on pH of the HAs solution. HAs obtained from fallow (A,B) and grassland (C,D) amended with BioC in 1st and 28th month of field experiment, as well as HAs obtained from BioC.

Full size image

Influence of BioC amendment on structure and chemical properties of HAs in fallow and grassland: spectroscopic approach
The analyses of the HAs isolated from fallow and grassland amended with BioC showed changes in the structural properties of these compounds. The E2/6 parameter estimated from UV–Vis data was changing both under the influence of different BioC doses and during the 3 years of the experiment. However, it should be assumed that the observed changes were of a different nature for fallow (Fig. 5A) and for grassland (Fig. 5B), due to varied trends in the activity of BioC on the analysed soils.
Figure 5

Changes in E2/6 values obtained for HAs of fallow (A) and grassland (B) amended with BioC (0, 1, 2, 3 kg m−2) as a function of time. Average values from 3 replicates in each term, ± standard deviation. Other letter designations indicate significant differences between values at α  More

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Plant and insect materials
A total of 56 apple plants were grown from seeds and sampled for this study. Cultivated apple plants resulting from crosses between various cultivated apple varieties were used (M. domestica, referred to as “Dom”, N = 14, Table S1). The seeds were kindly provided by INRAE IRHS Angers that performed every year crosses for apple breeding programs. A total of 42 M. sylvestris plants were grown from field-collected seeds. These wild apple seeds originated from three out of the five known European wild apple populations (referred to as Danish: Syl_Dk, French: Syl_Fr and Romanian: Syl_Ro, N = 14 per population). Each population was represented by a single sampling site, and within each site, each seed was sampled on a single mother tree, so that each seedling has a different parental origin. Though M. domestica is usually grafted, new plants were grown from seed to eliminate the rootstock effect.
After field sampling, seeds were stored at -20 °C before vernalization for the experiment. Seeds were then vernalized for three months at 4 °C in the dark, then grown in controlled conditions for two months before being individually transferred to 3 L pots containing commercial sterilized potting soil. Potted plants were grown in a growth chamber for four weeks under the following conditions: 20 ± 1 °C, 75 ± 5% Relative Humidity (RH), and a 16:8 light:dark (L:D) photoperiod. The 56 plants were then genotyped using 13 previously published microsatellite markers (see below) to confirm their genetic status (i.e., belonging to one of the M. sylvestris European populations or crop-to-wild/wild-to-wild hybrid).
A single colony of D. plantaginea (Hemiptera: Aphididae) was used and provided by INRAE which were sampled as a population in spring 2018 from an apple tree at the Agrocampus Ouest orchard (Angers, France) (Philippe Robert, personal communication). This aphid population was mass reared without differentiating individual aphid clones on M. domestica cv. “Jonagold” plants obtained by in vitro multiplication21. Pots containing three plants (90 × 90 × 70 mm) were placed in a Plexiglas cube (50 cm). Mass rearing and experiments were performed in growth chambers under 20 ± 1 °C, 60 ± 5% RH, and a 16:8 L:D cycle.
Synchronized first instar nymphs were obtained by placing parthenogenetic adult females on plantlets for 24 h before removing them. They were then reared on M. domestica cv. “Jonagold” plants inside Plexiglas aerated boxes (36 × 24 × 14 cm) for ten days then used as the young adult RAA for the behavioral/performance experiments.
Apple population genetic diversity and structure
Genomic DNA was extracted with the NucleoSpin plant DNA extraction kit II (Macherey & Nagel, Düren, Germany) according to the manufacturer’s instructions. Microsatellites were amplified by multiplex PCR, with the Multiplex PCR Kit (QIAGEN, Inc.). We used 13 microsatellite markers, Ch01f02, Ch01f03, Ch01h01, Ch01h10, Ch02c06, Ch02c09, Ch02c11, Ch02d08, Ch03d07, Ch04c07, Ch05f06, GD12, and Hi02c07 in four multiplexes (MP01, MP02, MP03, MP04)4. PCR were performed in a final reaction volume of 15 ml (7.5 ml of QIAGEN Multiplex Master Mix, 10–20 mM of each primer, with the forward primer labelled with a fluorescent dye and 10 ng of template DNA) (See4 for more details). The final volume was achieved with distilled water. A touch-down PCR program (initial annealing temperature of 60 °C, decreasing by 1 °C per cycle down to 55 °C) was used. Genotyping was performed on the GENTYANE platform (INRAE Clermont-Ferrand) using an ABI PRISM X3730XL, with 2 ml of GS500LIZ size standard (Applied Biosystems). Alleles were scored with GENEMAPPER 4.0 software (Applied Biosystems). Only multilocus genotypes with  0.1 were classified as crop-to-wild hybrids (i.e., introgressed by M. domestica). Once crop-wild hybrids removed, plants assigned to a given wild gene pool with a cumulated membership coefficient  > 0.9 were defined as “pure wild” individuals. Plants assigned to the wild gene pool with a cumulated membership coefficient  More

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